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1 Biological Science and Technology, Tokai University, Numazu, Shizuoka, Japan
2 Physiology and Biophysics, University of Calgary, Calgary, Alberta, Canada
* To whom correspondence should be addressed. E-mail: manabu{at}tokai.ac.jp.
Lymnaea can be classically conditioned pairing photic stimulation with a rotational stimulus. The electrophysiological properties of the Lymnaea photoreceptors and statocyst neurons are incompletely known. There are two types of ocular photoreceptors and three types of statocyst 'hair-cells'. Type A photoreceptor had a response latency from 400 to 200 ms, with a graded depolarizing response having maximum action spectra at 480-500 nm, corresponding to the 
max of rhodopsin. Additionally they extend their axons in the direction of the other type of photoreceptor neuron, the Type T cell. These neurons have a two-component response to light: a response reversibly reduced in Ca2+-free saline, and a component persisting in Ca2+-free saline. Type T cells send processes into the cerebral ganglion and terminate close to the ending of the statocyst hair cells. Hair cells send their terminal branches to the cerebral ganglia close to the terminations of the T Type cells. Caudal hair cells respond to a light flash with a depolarization, while the rostral cells respond with a hyperpolarization. The response latency in all hair cells was dependent on the stimulus intensity; the brightest light tested had a latency of 200 ms. The photo-induced response was abolished in Ca2+-free saline, while it was still present in high Ca2+- high Mg2+ saline, consistent with the hypothesis that the connection between the photoreceptors and hair cells is monosynaptic. Thus, the sensory information necessary for forming an association between photic and rotational stimuli converges on the statocyst neurons.
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